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Mesozoic Fishes 4 - Homology and Phylogeny, G. Arratia, H.-P. Schultze & M. V. H. Wilson (eds.): pp. 433-442, 2 figs.
©2008 by Verlag Dr. Friedrich Pfeil, Miinchen, Germany - ISBN 978-3-89937-080-5

A review of the Mesozoic Record
of the Carcharhiniformes
Charlie J. UNDERWOOD & David J. WARD

Abstract
Although the Carcharhiniformes represent one of the most diverse and important groups of sharks alive today,
their early history is very poorly known. Reinterpretation of previously figured Jurassic and Cretaceous fossils,
along with collection of new specimens from the Cretaceous of the British Isles, has allowed the early record
of this order to be reinterpreted. Whereas members of only one carcharhiniform family have been previously
recorded from the Jurassic, and two from the Cretaceous, it is considered here that fossils of two families are
known from Jurassic rocks, and at least five families from the Cretaceous. The relative timing of familial appearances is consistent with the predictions derived from cladistic analyses, although some of the cladogenic events
can now be shown to have been earlier than previously recognised.
Introduction
The Carcharhiniformes sensu COMPAGNO (1973) represent one of the most diverse groups of selachians.
Modern examples of this order vary in size from under 30 centimetres to over five metres in length, and
are distributed from the ocean floor to intertidal areas, with some species being oceanic pelagics and others entering fresh water. Although much of the radiation within carcharhiniform families occurred within
the Caenozoic, a number of families first appeared within the Jurassic and Cretaceous. Despite their importance within modern marine environments and their phylogenetically predicted early radiation (e.g.,
SHIRAI 1996), the early record of the group is very poorly known. There are eight extant families: the
Carcharhinidae, Hemigaleidae, Leptochariidae, Proscylliidae, Pseudotriakidae, Scyliorhinidae, Sphyrnidae,
and Triakidae. Only the Scyliorhinidae and the Triakidae have previously been recognised in the Mesozoic,
although a number of carcharhiniform teeth have been figured that do not closely match those of extant
members of those families (UNDERWOOD 2006).
Material
Much of the material considered here has been figured in other publications, although the taxonomic affinities of a number of these specimens have been reassessed for this study as described in the text below. In
addition, a number of taxa are considered here that have not yet been figured elsewhere. A number these
taxa form part of extensive faunas currently under study by the authors. These specimens were recovered
from a number of horizons of phosphatic chalk from the Santonian to Lower Campanian of southern England, and from a Coniacian phosphatic greensand from Northern Ireland. In addition, a triakid tooth from
the British Cenomanian found in the collections of the Natural History Museum, London, is figured.
All specimens figured here are deposited in the Natural History Museum, London (prefix BHNM P.)
or in the Liverpool Museum (National Museums and Galleries on Merseyside) (prefix LIVCM).

433

Jurassic Carcharhiniformes
The earliest records of Carcharhiniformes are from the Bathonian, where several genera are recognised
(e.g., UNDERWOOD & W A R D 2004, CAPPETTA, pers. comm. 2004). Earlier reported records are now
known to have been based on misidentified teeth of palaeospinacid sharks (DELSATE et al. 2002). Some
of these earliest records represent members of the Scyliorhinidae GILL, 1862 (represented today by the
catsharks or spotted dogfish), including the taxa Palaeoscyllium tenuidens UNDERWOOD & WARD, 2004
(Fig. 1 A) and an additional unnamed form from the Bathonian of England. The familial position of other
taxa is less clear. The genera PraeproscyIlium UNDERWOOD & WARD, 2004 (Fig. 1E,F) and Eypea UNDERWOOD & WARD, 2004 (Fig. 1G) both have dental morphologies, with regard to the form of both the
crown and the root, closer to extant members of the Proscylliidae FOWLER, 1941 (finback catsharks) than
Scyliorhinidae, and are here considered to be probable members of that family. Fragmentary teeth of an
additional, larger, ?carcharhiniform lack diagnostic morphological characters.
Carcharhiniform material from Late Jurassic rocks suggests that taxa similar to those living in the Bathonian were present. Palaeoscyllium WAGNER, 1857 is known from both skeletal and dental remains (see
LEIDNER & THIES1999, UNDERWOOD 2002). The shape of the body and head, position and shape of the
fins, morphology and heterodonty of the teeth, and shape of the placoid scales all more closely resemble
those of extant scyliorhinids than members of other carcharhiniform families, agreeing with a scyliorhinid
affinity for the genus. Other Late Jurassic taxa are far less well known, but suggest the presence of several
genera. Unnamed teeth from the Kimmeridgian of France (CANDONI 1995) appear to belong to a species
of Praeproscyllium (UNDERWOOD 2004) or an allied genus. A single tooth from the Kimmeridgian of
Portugal (KRIWET 1997) appears to show some similarity to Praeproscyllium, but may represent a further,
unnamed, genus. Poorly preserved skeletal remains of Macrourogaleus hassei (WOODWARD 1889) appear
to represent a scyliorhinid with a single dorsal fin (KRIWET & KLUG 2004). A well-preserved skeleton
of what is probably an additional Tithonian carcharhiniform taxon is currently under study (THIES pers.
com. 2003). Although sometimes regarded as a carcharhiniform (THIES & CANDONI 1998), we consider
Corysodon SAINT-SEINE, 1949 to be an orectolobiform based on the morphology of the tooth root and the
shape and positions of the fins.

Cretaceous Carcharhiniformes
Teeth of small carcharhiniform sharks are frequently abundant and diverse elements within Cretaceous
marine shark assemblages, with high diversity assemblages being present within many Late Cretaceous
assemblages (e.g., CAPPETTA 1980, NOUBHANI & CAPPETTA 1997, and several samples currently
under study by the authors). In addition, there are several known occurrences of well-preserved skeletal
remains, particularly from the Upper Cretaceous lithographic limestones of Lebanon (CAPPETTA 1980),
which in many cases confirm the affinities of the isolated teeth. Despite this, the very small size of many
of the isolated teeth has resulted in them commonly being overlooked. The lack of detailed study of the
Fig. 1.
Teeth of representative Jurassic and Cretaceous Carcharhiniformes. White scale bars for all species = 500 pm. >
All photographs SEM images except Q. A, Palaeoscyllium tenuidens UNDERWOOD & WARD, 2004, BMNH
P. 66045, labial view of lateral tooth, Bathonian of Watton Cliff, Dorset. B, Palaeoscyllium sp., BMNH P. 66293,
labial view of anterior tooth, fluvial Barremian of Yaverland, Isle of Wight. C,D, Pseudoscyliorhinus sp., BMNH
P. 66366, labial (C) and lingual (D) views of lateral tooth. Santonian of Boxford, Berkshire. E, Praeproscyllium
oxoniensis UNDERWOOD & WARD, 2004, BMNH P. 66054, labial view of anterior tooth, Bathonian of Woodeaton Quarry, Oxfordshire. F, Praeproscyllium oxoniensis UNDERWOOD & WARD, 2004, BMNH P. 66055, labial
view of posterior tooth, Bathonian of Woodeaton Quarry, Oxfordshire. G, Eypea leesi UNDERWOOD & WARD,
2004., BMNH P. 66058, labial view of anterior tooth, Bathonian of Watton Cliff, Dorset. H,I, Pteroscyllium sp.,
BMNH P. 66395, lingual (I) and labial (H) views of lateral tooth. Santonian of Winterbourne, Berkshire. J,K, Leptocharias sp., BMNH P. 66418, labial (J) and lingual (K) views of tooth. Santonian of Winterbourne, Berkshire.
L,M, Leptocharias sp., BMNH P. 66420, labial (L) and lingual (M) views of tooth. Santonian of Winterbourne,
Berkshire. N,0, ?triakidae indet., LIVCM 1998.48.E, labial (O) and lingual (N) views of tooth. Hauterivian of
Speeton, Yorkshire. P, Paratriakis sp. BMNH P. 66415, Santonian of Winterbourne, Berkshire. Q, Pachygaleus sp.,
BMNH P. 13346, labial view of tooth. Cenomanian of southern England. R,S,T, Ihoxodon sp., BMNH P. 66422,
occlusal (R), labial (S) and lingual (T) views of tooth. Santonian of Winterbourne, Berkshire.
434

435

I

isolated dental remains of Cretaceous Carcharhiniformes has resulted in a very poor understanding of
the generic, and probably familial, diversity.
Early Cretaceous marine selachian faunas are generally poorly known, with only Albian faunas being relatively well represented in the literature. Despite this, scattered records suggest the presence of a
number of taxa in Berriasian to Aptian rocks. The Scyliorhinidae are represented by PalaeoscyIlium, which
has been recorded from the Valanginian (REES 2005), Barremian (SWEETMAN & UNDERWOOD 2006)
(Fig. IB), and Albian (UNDERWOOD & MITCHELL 1999). Cretascyliorhinus UNDERWOOD & MITCHELL,
1999 is known from a number of Albian sites (e.g., CAPPETTA 1977), as are a number of scyliorhinids of
uncertain generic affinity (e.g. UNDERWOOD & MITCHELL 1999, W A R D in press). Additional Valanginian scyliorhinids are mentioned, but not described or figured (CAPPETTA 1990: 35).
Protoscyliorhinus HERMAN, 1977 has been recorded from a number of Early Cretaceous sites, ranging
from Barremian (BIDDLE & LANDEMAINE 1988) to Albian (BIDDLE 1993) in age. The dental morphology
of Protoscyliorhinus, with a "U"-shaped root lacking a clearly defined basal face, lacking clearly defined
lateral cusplets, and having a flat labial surface, is here considered to more closely resemble that of extant
members of the Proscylliidae than Scyliorhinidae, and so it is here tentatively considered to represent a
member of the former. A second genus of uncertain affinity, Pteroscyllium CAPPETTA, 1980 is known from
the English Aptian (UNDERWOOD 2004) and Albian (UNDERWOOD & MITCHELL 1999). Santonian examples of this genus are known from well-preserved skeletal remains from the Lebanon (CAPPETTA 1980)
that demonstrate a scyliorhinid-like overall morphology and dentition. However, the dental morphology
is unlike that of any known scyliorhinid (UNDERWOOD 2004), and more closely resembles that of some
more primitive taxa of Lamniformes. It is therefore considered here that this genus probably requires the
erection of a new family, although it is considered that the definition of this would be premature without
study of the skeletal remains.
A single tooth recorded from the Hauterivian of northern England appears to suggest the presence of
a species belonging to the Triakidae GRAY, 1851 (the family that includes the tope and smoothhounds)
(UNDERWOOD et al. 1999: 292) (Fig. 1N,0). The crown and root morphology is extremely similar to that
of many modern triakids and unlike that of anything else known from the early Cretaceous. The histology
could not be observed, and more material is needed to confirm the affinity of this taxon.
Although commonly abundant and diverse, much work is needed before the generic diversity of Late
Cretaceous scyliorhinids can be realistically assessed. Scyliorhinus BLAINVILLE, 1816 s.s. is known from
the Campanian and Maastrichtian (HALTER 1994), but the majority of species referred to Scyliorhinus lack
the distinctive dental characteristics of extant members of the genus (UNDERWOOD 2006). Palaeoscyllium
is present within the British Santonian and Campanian (pers. observ.) and Cretascyliorhinus is known
from a number of Upper Cretaceous stratigraphical intervals (see UNDERWOOD & MITCHELL 1999).
Pseudoscyliorhinus MULLER & DIEDRICH, 1991 is known from the Cenomanian (MULLER & DIEDRICH
1991), Turonian (as 'Scyliorhinus' reussi HERMAN, 1977), Coniacian, Santonian, and Campanian (pers. obs.)
(Fig. 1C,D). Some Late Cretaceous scyliorhinid species have tooth morphologies resembling extant genera
and may prove to be congeneric. Other species have a tooth morphology unlike any living scyliorhinid
species and will almost certainly require the erection of new genera.
Protoscyliorhinus, and hence possible representatives of the Proscylliidae, is recorded from the Cenomanian of Lithuania and Turonian of France (HERMAN 1977: 259). This is very similar in overall tooth
morphology to the Palaeogene genus Foumtizia NOUBHANI & CAPPETTA, 1997, which likewise may be
a proscylliid.
A number of species of Pteroscyllium have been recorded from the Late Cretaceous (Fig. 1H,I), with
the last records of the genus being from the Maastrichtian (NOUBHANI & CAPPETTA 1997). Skeletal
remains of two species are known from the Santonian of Lebanon (CAPPETTA 1980), and the body outline
and dentition are well known. The well-developed covering of denticles obscures much of the cranial and
postcranial skeleton, and these are therefore not so well known.
Although there is no recorded fossil record of the Leptochariidae GRAY, 1851 (represented today by
a single species, the barbeled houndshark), it is here considered to be represented within Late Cretaceous
assemblages. Isolated teeth recovered from several British Santonian sites (Fig. 1J-M) have a crown and
characteristic root morphology of Leptocharias Smith in MULLER & HENLE, 1838. It is considered here that
other unrecognised occurrences of this genus also occur in the fossil record, with teeth of the Palaeogene
'Scyliorhinus' ptychtus NOUBHANI & CAPPETTA, 1997 also strongly resembling Leptocharias and here
provisionally assigned to it.

436

Pn eproscyllium
Eypea
Unnamed genusUnnamed genus^
Macrourogaleus (:+? unr amed genus)
Protoscyliorhinus
Foumtizia
P'alaeoscyllium
Un named genus
probable unn amed genera
Cretascyliorhinus various unnamed generaPseudoscyliorhinus
Scylidrhinus
Casieria
Microscyliorhinus
Porodermoides
Prerriontreia
Pteroscyllium -

Pteroscyllium
genus indet.-

Pachygaleus
Paratriakisr
Palaeogaleus ArchaedtriakisSquatigaleus Triakis
Khouribgaleus
Galeorhinus
?

LoxodonPhysogaleus
Abdounia
Danogaleus

Carcharhinidae:
Leptocharias

Leptochariidae
Aal

Baj

Bat

M. Jurassic

Cal

Oxf

Kim

L. Jurassic

Tit

Ber

Val

Hau

Ber

Apt

Early Cretaceous

Alb

Cen

Tur

Con

San

Cam

Maa

Late Cretaceous

Fig. 2.
Known Mesozoic and Palaeocene ranges for carcharhiniform genera, including unnamed genera mentioned in the text. Note that the apparent Palaeocene radiation may be an artefact of the intensive study of material of this age from Moroccan phosphorites.

Triakids are typically common within Late Cretaceous assemblages, with well-preserved skeletons
being known from several European sites (e.g., CAPPETTA 1980, MULLER 1989). Material referred to
the extant genus Galeorhinus BLAINVILLE, 1816 is known from a number of species ranging from the
Cenomanian (Popov & Lapkin 1999, pers. observ., see Fig. 1Q) onWARDs. Despite this, the tooth morphology of Cretaceous nominal Galeorhinus species (teeth robust with a convex anterior edge, short main cusp
and swollen or folded labial crown edge) suggest that these species probably represent Pachygaleus CAPPETTA, 1992. An additional, unnamed triakid is also present in the Cenomanian of Germany (MULLER
& DIEDRICH 1991). Paratriakis HERMAN, 1977 (Fig. IP) first appears in the latest part of the Cenomanian
(HERMAN 1977). Within the latter parts of the Late Cretaceous, other genera appear alongside Pachygaleus
and Paratriakis. Palaeogaleus GURR, 1962 is first recorded in the Santonian (pers. observ.), and continues
into the Palaeogene. Withirt the Campanian and Maastrichtian, Archaeotriakis CASE, 1978 and Squatigaleus
CAPPETTA, 1989 are recorded respectively. These both have highly derived dental morphologies and may
be closely related. Their teeth to some extent resemble the extant Pseudotriakis BRITO CAPELLO, 1868, but
the triakid affinity suggested by CAPPETTA (1989) is followed here.
The Carcharhinidae JORDAN & EVERM ANN, 1896 (here taken to include genera such as Rhizoprionodon
WHITLEY, 1929 and Loxodon MULLER & HENLE, 1838) comprise probably the most morphologically and
ecologically diverse family of modern sharks, including the whaler or requiem sharks. Despite this, much
of the radiation within this group appeared to occur during the Neogene (e.g., CAPPETTA 1987), and
the recorded fossil record does not extend earlier than the Palaeocene (e.g., MAISEY et al. 2004). Samples
from the Santonian currently under study by the authors, however, yielded rare teeth assignable to Loxodon (Fig. 1R-T) (UNDERWOOD 2006). This would represent the first known Mesozoic occurrence of this
family. Although dentally very similar to the genera Rhizoprionodon and Scoliodon MULLER & HENLE,
1837, these teeth have been provisionally assigned to Loxodon on the basis of crown morphology and lack
of an incipient posterior cusplet (HERMAN et al. 1991; pers. observ. of modern dentitions).
Environmental palaeoecology
Modern Carcharhiniformes are present within almost all marine environments (e.g., COMPAGNO 1988),
with some entering freshwater. Morphological diversity is highest in shelfal settings, with both large and
small taxa commonly being present. Some deeper water environments have a high species diversity, but
these are typically dominated by small, morphologically conservative forms (e.g., LAST & STEVENS
1994). Within the Neogene to Recent, medium to large (two metre plus) species are known within several
carcharhiniform families (Carcharhinidae, Hemigaleidae, Pseudotriakidae, and Sphyrnidae). Despite this,
there is no evidence for any Mesozoic carcharhiniforms of this size.
Although palaeoenvironmental coverage of published selachian faunas is very incomplete, fossil Carcharhiniformes have been recorded from a wide range of shelfal palaeoenvironments. The almost complete
lack of studies on sharks from Mesozoic deep oceanic facies results in a general lack of information on deep
water taxa. Middle Jurassic Carcharhiniformes are recorded from a number of different palaeoenvironments
(UNDERWOOD & W A R D 2004), with different taxa being recorded from different facies. Within the study
area, Eypea dominated offshore settings, with Palaeoscyllium in shallow water carbonates and Praeproscyllium in lagoons. This indicates that, even at this early stage in the evolution of the order, considerable
environmental specificity was present. Although all of these palaeoenvironments were marine to some
degree, molluscan faunas from some of the lagoonal deposits suggests reduced salinity and indicates the
ability of at least this genus of early carcharhiniform to survive in brackish conditions. Other Jurassic taxa
are known from neritic mudstones or lagoonal plattenkalks, with Palaeoscyllium formosum being present
in both, although it is probable that many of the plattenkalk faunas are allochthonous (VIOHL 1996), or
at least parautochthonous and derived from a range of surrounding habitats.
The majority of Early Cretaceous Carcharhiniformes are known from offshore marine sediments. Despite this, rare examples of scyliorhinid teeth are known from freshwater facies. Teeth of Palaeoscyllium
were recorded from a channel sandstone within the fluvial Barremian Wessex Formation of southern
England (SWEETMAN & UNDERWOOD 2006). It is unclear whether this represents a euryhaline or truly
freshwater species, but it is assumed, by analogy with modern carcharhiniforms, that the former is more
likely. No other species of the Scyliorhinidae, fossil or extant, is known to enter freshwater.
Although Late Cretaceous Carcharhiniformes are known from many sites; the great majority of these
are within offshore, fine grained (mudstone and chalk) facies. A small number of taxa, mostly triakids, are
438

known from shallow water sands and silts. Diversity is highly variable between sites, typically being low
within mudstones and chalks of the American Interior Seaway and higher within the northern European
Chalk facies. Diversity is highest within phosphatic deposits, such as from the southern margin of Tethys
(e.g., NOUBH ANI & CAPPETTA 1997). Even more diverse carcharhiniform assemblages are present within
the phosphatic chalks of the British Santonian and Campanian (with up to 12 carcharhiniform species in
some samples, currently under study by the authors), where the Carcharhiniformes form part of a very
diverse assemblage of small-toothed sharks, with over 90 % of teeth recovered being under 3 mm high.
Phylogenetic implications
Both morphological and molecular data have been used to reconstruct neoselachian phylogenies, and these
studies have typically included several Carcharhiniforme taxa. An analysis of interrelationships within the
Carcharhiniformes by C O M P A G N O (1988) included a greater variety of taxa, but resolution within the
group was poor. In all phylogenies, the Scyliorhinidae are placed as a sister group to all other (included)
Carcharhiniforme families (e.g, SHIRAI 1996, MAISEY et al. 2004, WINCHELL et al. 2004) or at least
close to the base of the clade (COMPAGNO 1988). WINCHELL et al. (2004) included two genera of the
Scyliorhinidae and concluded that the family is paraphyletic, with Apristurus GARMAN, 1913 being more
closely related to other carcharhiniform families than Scyliorhinus. The Proscylliidae are also considered
to be close to the base of the carcharhiniform clade, and are generally considered more derived than the
Scyliorhinidae (e.g., SHIRAI 1996, MAISEY et al. 2004). They appear to be close to, and possibly form a
monophyletic group with, Pseudotriakis (COMPAGNO 1988, SHIRAI 1996).
The relative positions of Leptocharias and the Triakidae are uncertain (SHIRAI 1996), and it is probable
that the latter is paraphyletic (MAISEY 1984, COMPAGNO 1988, SHIRAI 1996). The Leptochariidae and
Triakidae together form paraphyletic sister groups to the more derived clade containing the Carcharhinidae,
Sphyrnidae GILL, 1862 and Hemigaleidae HASSE 1879 (e.g., C O M P A G N O 1988, SHIRAI 1996, MAISEY
et al. 2004, WINCHELL et al. 2004). The Hemigaleidae appear to be a sister group to the Carcharhinidae
(including Sphyrnidae) (e.g. SHIRAI 1996), with the Carcharhinidae being paraphyletic if the Sphyrnidae are
excluded and Rhizoprionodon and Galeocerdo MULLER & HENLE, 1838 included (e.g., NAYLOR 1992).
The Middle Jurassic occurrences of scyliorhinids and probable proscylliids agree well with these
groups being basal within the carcharhiniform clade. The absence of pseudotriakids from the Jurassic (and
subsequent) fossil record may either be due to a consistently deep water habit of the family, or possibly
due to a relatively recent derivation of the Pseudotriakidae from the Proscylliidae.
The Cretaceous fossil record of the Triakidae and probable Leptochariidae again fits well with the
sequence of familial originations inferred from phylogenetic studies, although the poor Early Cretaceous
neoselachian record and the difficulty in recognition of leptochariids on dental remains suggests that there
is a good possibility of earlier specimens being recorded in the future.
The presence of Loxodon in the Cretaceous suggests a far earlier radiation of the 'crown group carcharhinids' than previously realised, as this is generally considered to be more derived than the Hemigaleidae.
Although none of these other groups are currently known from the Cretaceous, examples of Hemigaleidae,
Carcharhinidae and Galeocerdo are known from the Palaeocene and Eocene (CAPPETTA 1987, MAISEY et
al. 2004).
A summary of the known ranges of Mesozoic carchariniform genera is given in Fig. 2.
Acknowledgements
We would like to thank Andy GALE, Ian JARVIS and Peter WOODROOF for his help in the original collecting of
the British Late Cretaceous material considered here. Steven SWEETMAN is thanked for bringing our attention
to the Wessex Formation material. Alison LONGBOTTOM and staff at the Natural History Museum, London and
thanked for their help. We would also like to thank the referees whose comments greatly improved this paper.
This work was partly carried out with the assistance of University of London grant GL2CU.

439

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C l M < > ^

M

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WOODWARD, A. (1889): Catalogue of the fossil Fishes in the British Museum. 474 pp.; Part 1: London (British
Museum, Natural History)

Authors' addresses:
Charlie J. UNDERWOOD, School of Earth Sciences, Birkbeck College, Malet Street, London WC1E7HX, United
Kingdom; e-mail: c.Underwood@bbk.ac.uk
David J. WARD, Crofton Court, 81 Crofton Lane, Orpington, Kent, BR51HB, United Kingdom;
e-mail: david@fossil.ws.
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